Genetic variation in the endangered <Emphasis Type="Italic">Anisodus tanguticus</Emphasis> (Solanaceae), an alpine perennial endemic to the Qinghai-Tibetan Plateau

We used random amplified polymorphic DNA markers (RAPDs) to assess genetic variation between- and within-populations of Anisodus tanguticus (Solanaceae), an endangered perennial endemic to the Qinghai-Tibetan Plateau with important medicinal value. We recorded a total of 92 amplified bands, using 12 RAPD primers, 76 of which (P = 82.61%) were polymorphic, and calculated values of H_t and H_sp of 0.3015 and 0.4459, respectively, suggesting a remarkably high rate of genetic variation at the species level. The average within-population diversity also appeared to be high, with P, H_e and H_pop values of 55.11%, 0.1948 and 0.2918, respectively. Analyses of molecular variance (AMOVA) showed that among- and between-population genetic variation accounted for 67.02% and 32.98% of the total genetic variation, respectively. In addition, Nei’s coefficient of differentiation (G_ST) was found to be high (0.35), confirming the relatively high level of genetic differentiation among the populations. These differentiation coefficients are higher than mean corresponding coefficients for outbreeding species, but lower than reported coefficients for some rare species from this region. The genetic structure of A. tanguticus has probably been shaped by its breeding attributes, biogeographic history and human impact due to collection for medicinal purposes. The observed genetic variations suggest that as many populations as possible should be considered in any planned in situ or ex situ conservation programs for this species.     

Allozyme diversity in the federally threatened golden paintbrush, <Emphasis Type="Italic">Castilleja levisecta</Emphasis> (Scrophulariaceae)

Castilleja levisecta (Scrophulariaceae), the golden paintbrush, is an insect-pollinated herbaceaous perennial found in the Pacific Northwest. Currently restricted to two island populations off British Columbia and nine populations (eight on islands) in Washington, C. levisecta is a rare species threatened with extinction. Allozymes were used to describe genetic diversity and structure in these eleven populations. Despite its threatened status and small geographic range, exceptionally high levels of genetic diversity are maintained within C. levisecta. All sixteen of the loci resolved were polymorphic within the species (P_s=100%), while the mean percentage of loci polymorphic within populations (P_p) was 65.7%. The mean number of alleles per polymorphic locus (AP_s) was 2.94 within the species and averaged 2.38 within populations (AP_p). Genetic diversity (H_es) was 0.285 for the species, whereas mean population genetic diversity (H_ep) was 0.213. Smaller populations had, on average, fewer observed alleles and less genetic diversity. A significant negative correlation (r = –0.72) was found between genetic identity and geographic distance, indicating reduced gene flow between distant populations. The most geographically isolated population was one of the larger populations, one of the most genetically diverse and the most genetically divergent. A wide range of pairwise population genetic identities (I = 0.771 – 0.992) was found, indicating considerable genetic divergence between some populations. Overall, 19% of the total genetic diversity was distributed among populations. Results of this survey indicate that genetic augmentation of existing populations is unnecessary. The high allelic diversity found for the species and within its populations holds promise for conservation and restoration efforts to save this rare and threatened plant species.     

Genetic variation and clonal structure of the rare,riparian shrub <Emphasis Type="Italic">Spiraea virginiana</Emphasis> (Rosaceae)

Genetic diversity is often considered important for species that inhabit highly disturbed environments to allow for adaptation. Many variables affect levels of genetic variation; however, the two most influential variables are population size and type of reproduction. When analyzed separately, both small population size and asexual reproduction can lead to reductions in genetic variation, although the exact nature of which can be contrasting. Genetic variables such as allelic richness, heterozygosity, inbreeding coefficient, and population differentiation have opposite predictions depending upon the trait (rarity or clonality) examined. The goal of this study was to quantify genetic variation and population differentiation in a species that resides in a highly stochastic environment and is both rare and highly clonal, Spiraea virginiana, and to determine if one trait is more influential genetically than the other. From populations sampled throughout the natural range of S. virginiana, we used microsatellite loci to estimate overall genetic variation. We also calculated clonal structure within populations, which included genotypic richness, evenness, and diversity. Gene flow was investigated by quantifying the relationship between genetic and geographic distances, and population differentiation (θ) among populations. Observed heterozygosity, genotypic richness, and inbreeding coefficients were found to be representative of high clonal reproduction (averaging 0.505, 0.1, and –0.356, respectively) and the number of alleles within populations was low (range = 2.0–3.6), being more indicative of rarity. Population differentiation (θ) among populations was high (average = 0.302) and there was no relationship between genetic and geographic distances. By examining a species that exhibits two traits that both can lead to reduced genetic variation, we may find an enhanced urgency for conservation. Accurate demographic counts of clonal species are rarely, if ever, possible and genetic exploration for every species is not feasible. Therefore, the conclusions in this study can be potentially extrapolated to other riparian, clonal shrubs that share similar biology as S. virginiana.     

Genetic variation of <Emphasis Type="Italic">Oplopanax elatus</Emphasis> (Nakai) Nakai (Araliaceae)

Using allozyme analysis, genetic and genotypic variation of rare relict species, a member of one of ancient angiosperm families, Araliaceae, the clonal plant Oplopanax elatus (Nakai) Nakai, was evaluated. Electrophoretic separation of the enzymes is described, and genetic interpretation of the enzymes variation patterns is presented. The values of genetic variation indices obtained were low (P = 25%; A = 1.45: H _o = 0.131; H _e = 0.113) and comparable with the data reported for the rare plant species and the representatives of the family Araliaceae. The main factors responsible for the polymorphism level observed might be the evolutionary history of the species and gene drift. The level of genotypic diversity (G/N = 0.76; D = 0.97) was substantially higher, compared to the values reported for species with vegetative reproduction (D = 0.62). These results suggest certain contribution of the propagation by seeds in the formation of the species of interest.     

Extremely Low Levels of Allozyme Variation in Southern Korean Populations of the Two Rare and Endangered Lithophytic or Epiphytic <Emphasis Type="Italic">Bulbophyllum Drymoglossum</Emphasis> and <Emphasis Type="Italic">Sarcanthus Scolopendrifolius</Emphasis> (Orchidaceae): Implications for Conservation

Allozyme variation was investigated in two local populations of Bulbophyllum drymoglossum and three populations Sarcanthus scolopendrifolius, two rare and endangered lithophytes and epiphytes from South Korea. Genetic diversity was extremely low within populations (mean H _e = 0.011 for B. drymoglossum; 0.002 for S. scolopendrifolius). Among the putative screened 21 loci, we found only one polymorphic locus for each species. Only one polymorphic locus, detected just one population of each species, revealed significantly high degree of population differentiation between and among populations (F _ST = 0.253 for B. drymoglossum and F _ST = 0.899 for S. scolopendrifolius). These results suggest that genetic drift (consequence of a very small effective population size), coupled with a limited gene flow would play a major role in shaping population genetics of these species in South Korea. The current status of both species (small population sizes, spatially isolated populations, and highly localized habitats) in addition to the extremely low levels of genetic diversity and reckless collection of endangered orchids by plant sellers, significantly threaten the long-term survival of these species in Korea. Conservation of the two species requires both in situ strategies, by introducing of genets to increase effective population sizes by minimizing adverse effects (e.g., outbreeding depression and genetic swamping by non-native genotypes), and ex situ strategies, such as collection of genets from clonal ramets.     

Determination of the population genetic structure of the invasive weed <Emphasis Type="Italic">Ageratina adenophora</Emphasis> using ISSR-PCR markers

A genetic analysis of nuclear DNA was performed by inter-simple sequence repeat-polymerase chain reaction (ISSR-PCR) technique on 32 populations of Ageratina adenophora, an invasive triploid weed in China. Among the 100 ISSR markers detected, 12 showed genetic variation both within and among the populations. Among the 446 amplified bands, 93.5% were found polymorphic. Most individuals (99%) displayed a unique ISSR fingerprint pattern, which yielded a high level of polymorphism (P _o = 93.5%) and genetic diversity (Nei’s H _T = 0.2354). The estimates of population variation, based on ISSR-PCR, were high, as measured by the analysis of molecular variance (AMOVA, F _ST = 0.3140), the Wright’s F-statistics (G _ST = 0.3453), and the Shannon’s information index (H _sp = 0.3716). AMOVA revealed 68.6% genetic variation within the populations and 91.2% within the provinces. The Mantel test showed that genetic distance was significantly correlated with geographic distance. Published in Russian in Fiziologiya Rastenii, 2009, Vol. 56, No. 3, pp. 453–459. This text was submitted by the authors in English.     

Conservation of long-lived perennial forest herbs in an urban context: <Emphasis Type="Italic">Primula elatior</Emphasis> as study case

Urban forests are generally fragmented in small isolated remnants, embedded in an inhospitable human-used matrix, and incur strong anthropogenic pressures (recreational activities, artificialization, pollution and eutrophication). These lead to particularly high constraints even for common forest herbs, whose genetic response may depend on life-history traits and population demographic status. This study investigated genetic variation and structure for 20 allozyme loci in 14 populations of Primula elatior, a self-incompatible long-lived perennial herb, occurring in forest fragments of Brussels urban zone (Belgium), in relation to population size and young plants recruitment rate. Urban populations of P. elatior were not genetically depauperate, but the small populations showed reduced allelic richness. Small populations showing high recruitment rates—and therefore potential rejuvenation—revealed lower genetic diversity (H _o and H _e) than those with low or no recruitment. No such pattern was observed for the large populations. There was a significant genetic differentiation among populations within forest fragments (F _SC = 0.052, P < 0.001), but not between fragments (F _CT = 0.002, P > 0.10). These findings suggest restricted gene flow among populations within fragments and local processes (genetic drift, inbreeding) affecting small populations, strengthened when there is recruitment. Urban forest populations of long-lived perennial herbs can be of conservation value. However, restoration of small populations by increasing population size through regeneration by seedling recruitment may lead to negative genetic consequences. Additional management, aiming to restore gene flow among populations, may need to be applied to compensate the loss of genetic diversity and to reduce inbreeding.     

Determining patterns of genetic diversity and post‐glacial recolonization of western Canada in the Iowa golden saxifrage,Chrysosplenium iowense (Saxifragaceae), using inter‐simple sequence repeats

Chrysosplenium iowense Rydb. (Saxifragaceae) is a southern Canadian boreal forest species with a small number of disjunct populations occurring in the Driftless Area of northeastern Iowa and southeastern Minnesota. This disjunction is attributed to the actions of glacial movement and climate change during the Pleistocene. Populations within each of these distributions may have been isolated for 115 000 years or more and, although levels of genetic divergence between these regions may be significant, there is no morphological or cytological variation associated with this geographic break. We employed inter‐simple sequence repeat (ISSR) markers to determine patterns of genetic diversity within 12 populations (six Canadian; six Iowan) of C. iowense and elucidate the routes of post‐glacial recolonization for the species. Despite finding relatively high levels of genetic divergence (θ^II = 0.383, θ^II = 0.299) between Driftless Area and Canadian populations, there is no conclusive evidence of a speciation event within C. iowense. Analyses show moderate levels of genetic diversity within the species (H_T = 0.188), the majority of which is partitioned among individuals within populations (68.18%), which were similar across the northern (H_T = 0.234, H_T = 0.28) and southern (H_T = 0.189) ranges. Finally, the patterns of genetic diversity within C. iowense suggest that the Canadian range was established by migrants originating in now extinct refugial populations that existed outside the Driftless Area. © 2008 The Linnean Society of London, Biological Journal of the Linnean Society, 2008, 95 , 815–823.     

Demography, and genetic and morphological variability of the endangered Sophronitis sincorana (Orchidaceae) in the Chapada Diamantina, Brazil

We carried out a demographic study and evaluated the genetic and morphological variability in five populations of the endangered Sophronitis sincorana (Orchidaceae) endemic to Northeastern Brazil, based on allozyme and morphometric analyses. Plant density was approximately 0.5 plants/m², and the projected total number of plants was approximately 50,000 individuals. However, fruit set and recruitment of individuals are rare. The genetic variability was very high in all populations (P = 100, A = 3.0−3.5, H _e = 0.33−0.48), and all populations presented similar values of morphological variability. Low genetic and morphological structuring were found in the species (F _ST = 0.053, A_MRPP = 0.018). The elevated coefficient of endogamy encountered in populations of S. sincorana indicates the occurrence of structuring within the populations. The lack of correlation between morphological and genetic variation in this species indicates that none of the markers examined should be used separately for either conservation purposes.     

Characterization of gene pools of three <Emphasis Type="Italic">Pinus pumila</Emphasis> (Pall.) Regel populations at the range margins

Genetic variation of Siberian dwarf pine Pinus pumila (Pall.) Regel was characterized in three marginal populations in southwestern, southern and eastern parts of the natural species range (Baikal Area, Primorye, Kamchatka) using isozyme analysis. Analysis involving 16 isozyme loci encoding ten enzyme systems was conducted. Our results confirm that P. pumila is one of the most polymorphic species in the genus Pinus. Three marginal populations exhibited high genetic variation (P ₉₅ = 68.8%, H _o = 0.247, H _e = 0.291). Populations heterogeneity and significantly high level of divergence in coniferous (F _ST = 0.050, D _N = 0.044) reflect their genetic originality. In summary, it was shown that the level of genetic variation characteristic for P. pumila in other parts of the not only is reproduced in the populations examined but even is close to maximum there.     

Genetic variability and differentiation of <Emphasis Type="Italic">Caragana microphylla</Emphasis> populations as revealed by RAPD markers

Genetic variability in random amplified polymorphic DNA (RAPD) was studied in 90 individuals of Caragana microphylla, an outcrossing perennial shrub species, from five natural populations sampled in Inner Mongolia steppe of China on a small scale. Nineteen selected primers were used to amplify DNA samples, and totally 225 bands were detected. The percentage of polymorphic bands within populations ranged form 58.22% to 63.56%, with an average of 60% at the population level and 71.11% at the species level, indicating relatively high genetic variations in C. microphylla species. Shannon’s information index (l) and Nei’s gene diversity (h) showed the similar trend with each other. According to the analysis of Nei’s gene diversity, the percentage of genetic variation among populations was 7.13%, indicating a low level of genetic differentiation among populations. There existed a strong gene flow (N _m = 3.26) among populations. Although AMOVA analysis also revealed most variation was within populations (Φ_ST = 4.1%), a significant proportion was observed among populations (P < 0.001) in the present study, suggesting genetic differentiation occurred among populations at a certain extent. Based on Mantel’s tests and the results of previous studies, the genetic structure pattern of C. microphylla accorded with the isolation-by-distance model on a very large scale, however, on a small scale, the significant genetic differentiation among populations might be enhanced by the micro-environmental divergence among the sampling sites, rather than by geographic factors. Analysis of the genetic variations of C. microphylla populations provided useful information for the adaptive strategy of Caragana species.     

Comparative analysis of genetic structure in natural populations of two <Emphasis Type="Italic">Arabidopsis</Emphasis> species with different degree of panmixia

Comparative analysis of genetic structure of northern natural populations of two Arabidopsis species with different degrees of panmixia was performed. The variability of 121 RAPD loci in seven populations of model plant A. thaliana possessing high degree of self fertility was studied together with 93 RAPD loci in population of cross-pollinating species A. lyrata ssp. petraea. The population of A. l. petraea demonstrated higher level of genetic variability (P _99% = 62.50%; H _exp = 0.169) than the populations of A. thaliana, which is obviously connected with biological features of reproduction of the species. A significant level of genetic variability (P _99% = 42.27%; H _exp = 0.126) was revealed in populations of A. thaliana, which is not typical for self-pollinating plant species. The high population polymorphism of A. thaliana in the northern part of its range may be connected with adverse environmental conditions. The genetic distances between populations of the species studied (average D _N = 0.494) confirm close relatedness between A. thaliana and A. l. petraea.     

Genetic structure of Polytrichum formosum in relation to the breeding system as revealed by microsatellites

Microsatellite variation was determined for three Danish and three Dutch populations of the haploid moss species Polytrichum formosum to gain insight into the relative importance of sexual vs. asexual reproduction for the amount and structure of genetic variation. In general, low levels of microsatellite variation were observed within this species. Even when estimated for polymorphic loci only, the levels of microsatellite variability (P=90.6, A=4.3 and H_S=0.468) within populations were on average lower than those reported for most other plant species. In contrast, genotypic diversity was high within each of the examined populations, indicating that sexual reproduction is a very important determinant of the genetic structure of P. formosum within populations. In agreement with previous findings for allozyme data, no significant genetic differentiation (F_ST=0.028, R_ST=0.015) was observed neither between populations nor between regions approximately 450 km apart (Denmark vs. the Netherlands). These low levels of population differentiation observed for both types of genetic markers are probably best explained by a high level of effective spore dispersal (gene flow) between populations. Therefore, also on a large geographical scale sexual reproduction is the most important determinant of the genetic structure of P. formosum, despite the high potential to reproduce clonally.     

A cline of allozyme variation inAbies mariesii

Genetic variation at 22 allozyme loci was examined for 1,003 trees from 11 isolated natural populations ofAbies mariesii covering all except the southernmost region of its geographic range. Genetic diversity within species (H _es=0.063) was low compared to many other long-lived woody species. Most of the genetic variation is found within populations (G _ST=0.144) despite their isolated distribution. Genetic distance between populations was positively correlated with geographic distance. Genetic diversity within populations was generally low (meanH _ep=0.054), but varied across populations in a clinal fashion such that genetic variation decreased with increasing latitude. These genetic characteristics may reflect the distribution history of this species.     

Population genetic structure of wild-growing ginseng (<Emphasis Type="Italic">Planax ginseng</Emphasis> C.A. Meyer) assessed using AFLP markers

Genetic variability in ten populations of wild-growing ginseng was assessed using AFLP markers with the application of fragment analysis on a genetic analyzer. The variation indices were high in the populations (P = 55.68%, H _S = 0.1891) and for the species (P = 99.65%; H _S = 0.2857). Considerable and statistically significant population differentiation was demonstrated (θ_B = 0.363; Bayesian approach, “full model”; F _ST = 0.36, AMOVA). The results of AMOVA and Bayesian analysis indicate that 64.46% of variability is found within the populations. Mantel test showed no correlation between the genetic and geographic distances among the populations (r = −0.174; p = 0.817). Hierarchical AMOVA and analysis of genetic relationships based on Euclidean distances (NJ, PCoA, and MST) identified two divergent population groups of ginseng. Low gene flow between these groups (N _m = 0.4) suggests their demographic independence. In accordance to the concept of evolutionary significant units (ESU), these population groups, in terms of the strategy and tactics for conservation and management of natural resources, should be treated as management units (MUs). The MS tree topology suggests recolonization of southern Sikhote-Alin by ginseng along two directions, from south and west.     

Genetic diversity,demographical history and conservation aspects of the endangered yew tree Taxus contorta (syn. Taxus fuana) in Pakistan

The western Himalayan yew (Taxus contorta Griffith, synonym Taxus fuana Nan Li & R.R. Mill), one of the highly prized tree species for its anticancer properties, is experiencing severe reduction in populations across its range in Pakistan. We examined a chloroplast DNA region (trnL-F) and ten nuclear microsatellite loci variations among seven populations to investigate the levels of genetic diversity and reconstruct the demographic history of T. contorta in Pakistan. A low haplotype diversity (H _T?=?0.182), moderate level of microsatellite diversity (H _E?=?0.541?±?0.034), significant population differentiation (F _ST?=?0.107) and high level of inbreeding (F _IS?=?0.219–0.418) characterized this species. Results of an approximate Bayesian computation (ABC), mismatch distribution analysis, neutrality tests and data on glacial chronologies of the western Himalaya indicated a historic demographic expansion of T. contorta populations from small ancestral populations. This expansion is estimated to have occurred at least 37.5 thousand years ago (kya) during the late Pleistocene. An unprecedented level of habitat disturbance over the last few decades, coupled with the highly variable climate may have played an important role in shaping the genetic structure of the extant T. contorta populations. We have discussed a range of sustainable management measures for the ecological restoration of remnant populations as well as for a sustainable exploitation of this plant of high economic potential.     

Does genetic variation and gene flow vary with rarity in obligate seeding Persoonia species (Proteaceae)?

Theory predicts that genetic variation is a determinant of persistence, and that the abundance and distribution of variation is strongly dependent on genetic drift and gene flow. Small, isolated populations are expected to be less diverse and more differentiated than large, inter-connected populations. Thus rare species may be more at risk of extinction. We used 389 putative AFLP loci to compare genetic variation and structuring in two pairs of closely-related common (large populations geographically widespread) and rare (small populations spatially restricted) Persoonia species. We genotyped 15–22 adult plants, from four populations, covering the geographic range of each species. Although genetic diversity was low for all four species (for long-lived outcrossing perennials), we found significantly more diversity within populations of the rare species than within those of the common species. AMOVA revealed significant levels of structure both among species (21%) and populations (15%). The proportion of inter-population variation within species did not vary consistently with rarity (Pair 1 rare 21.1% versus common 16.5%; Pair 2 rare 15.8% versus common 20.6%). However populations of the rare species were more differentiated than common species with similar geographic separation, suggesting greater gene flow between populations of the common species. Therefore, even relatively small genetically isolated populations of rare Persoonia species were more diverse than large populations of common Persoonia species. We hypothesise that common Persoonia species have undergone a rapid range expansion from a narrow gene pool, while genetic diversity is maintained in the soil seed-bank of rare remnants.     

Strong genetic differentiation of the East-Himalayan <Emphasis Type="Italic">Megacodon stylophorus</Emphasis> (Gentianaceae) detected by Inter-Simple Sequence Repeats (ISSR)

Megacodon stylophorus (Clarke) Smith is a perennial alpine herb endemic to the species-rich eastern Himalayan region. Its populations are locally scattered as isolated patches throughout this region. Genetic variation within and among six populations of this species was assessed using ISSR fingerprinting with 13 primers. High levels of genetic diversity exist within species (P = 69.83%, H_T = 0.1949 and H_sp = 0.3047), while the within-population diversity is low (P = 11.21%, H_E = 0.0532 and H_pop = 0.0792). Extraordinarily high levels of genetic differentiation were detected among populations based on various statistics, including Neis genetic diversity analysis (72.7%), Shannons diversity index (74.01%) and AMOVA (80.70%). That is, populations shared low levels of genetic identity (I = 0.8203 ± 0.0430). This genetic structure was probably due to severe genetic drift of the small-sized patchy populations resulting from postglacial habitat fragmentations. The observed genetic structure of the populations implies that as many populations as possible should be considered for any in situ and ex situ conservation practice on this species.     

Disparity of allozyme variation levels in three Magnolia (Magnoliaceae) species from the southeastern United States

Allozyme variation at 17 loci encoding ten enzyme systems was examined in 21, 22, and 15 populations across the ranges of Magnolia fraseri, M. macrophylla, and M. tripetala, respectively, in the southeastern United States. All three species have regional distributions, and are insect-pollinated outcrossing deciduous trees with seeds dispersed by birds, yet strikingly different levels of genetic variability were observed among them. In comparison with other woody angiosperm species, M. fraseri possesses a moderate amount of variation at the population level (A = 1.4, P = 39.2, H_o = 0.111, and H_e = 0.111), whereas M. macrophylla and M. tripetala are genetically depauperate in their populations (A = 1.2, P = 18.7, H_o = 0.47, and H_e = 0.055; A = 1.1, P = 11.0, H_o = 0.032, and H_e = 0.033). Examination of population structure revealed a small amount of inbreeding within populations and extensive intra- and interregional differentiation among populations of the latter two species. These two factors are perhaps partly responsible for the low genetic variability in populations of the two Magnolia species. Furthermore, the bottleneck effect caused by extinctions during cold periods of the Quaternary glaciations and human deforestation in the last two centuries as well as the founder effect in postglacial establishment of the populations might have also played significant roles in loss of genetic diversity in M. macrophylla and M. tripetala. We suggest that historical factors are important determinants of genetic variation profile of a species, in addition to life history and ecological characteristics as generally recognized.     

Low levels of genetic variation inPhenakospermum guyannense (Strelitziaceae), a widespread bat-pollinated Amazonian herb

Phenakospermum guyannense is a monotypic, arborescent, long-lived monocot that is widespread in Amazonian South America. This outcrossing species is pollinated primarily by phyllostomid bats. Given these life-history characteristics,P. guyannense is expected to exhibit high levels of genetic variation and gene flow. We used isozyme electrophoresis and randomly amplified polymorphic DNA (RAPD) to characterize genetic variation in populations ofP. guyannense from French Guiana. Both measures detected a surprisingly low level of genetic variation, with only five out of twenty (25%) allozyme loci polymorphic (P), 1.35 alleles per locus (A), and an expected heterozygosity (H_e) of 0.090 at the species level. Isozymic genetic variation was even lower within populations (P = 17.5, A = 1.24, H_e = 0.074), and was corroborated by a RAPD assay that used 26 arbitrary primers (P = 3.61, A = 1.04, H_e = 0.014). Although overall levels of variation were low, the detectable variation was distributed as would be expected for an outcrossing species with extensive gene flow (mean G_ST = 0.230). We suspect thatP. guyannense is depauperate in genetic variation because of a series of bottlenecks that affected the species over this portion of its range.